The Natural History of

edited by Philippe Bouchet, Hervé Le Guyader, Olivier Pascal

Patrimoines Naturels

The Natural History of

Santo

Publications scientifiques du muséum IRD ÉDITIONS P RO - NATURA INTERNATIONAL

The Santo 2006 expedition was organized, with the support, among others, of

and

Représentation de l'Union Européenne au Vanuatu


Santo

edited by Philippe Bouchet, Hervé Le Guyader & Olivier Pascal

Contents

Foreword by Edward Natapei, Prime Minister of Vanuatu ......................................    10

Introduction. The Natural History of Santo: An Attempt to Bridge the Gap between Academic Research and Consevation and Education  ...........................    11

Vanuatu in the South Pacific  .............................................................................................................    13 Benoît Antheaume

Espiritu santo in space and time ...........................................................................................    17 coordinated by Bruno Corbara

The Late Quaternaty Reefs .............................................................................................................................   19 Guy Cabioch & Frederick W. Taylor

The Holocene and Pleistocene Marine Faunas Reconsidered .................................   25 Pierre Lozouet, Alan Beu, Philippe Maestrati, Rufino Pineda & Jean-Louis Reyss

Geography of Santo and of the Sanma Province  ................................................................   34 Patricia Siméoni

Drainage, Hydrology and Fluvial Geomorphology .............................................................   46 James P. Terry

The Climate of Santo  ..............................................................................................................................................................   52 James P. Terry

Large-scale Climatic and Oceanic Conditions around Santo ...............................................................   57 Christophe Maes & David Varillon

A Brief History of Biodiversity Exploration and Scientific Expeditions on and off the Island of Santo  ......................................................................................................................................   62 Bruno Corbara & Bertrand Richer de Forges

Deforestation on Santo and Logging Operations ..........................................................................................   67 Rufino Pineda

The Impact of WWII on Infrastructures and Landscape  ..........................................................................   69 Laurent Palka & Rufino Pineda

Conservation Efforts in Santo .........................................................................................................................................   71 Samson Vilvil-Fare

vegetation and flora .....................................................................................................................................................    73 coordinated by Jérôme Munzinger & Porter P. Lowry II

Exploration by the Santo 2006 Botany Team  ....................................................................................................   75 Porter P. Lowry II & Jérôme Munzinger

Principal Types of Vegetation Occuring on Santo ..........................................................................................   76 Jérôme Munzinger, Porter P. Lowry II & Jean-Noël Labat

Phytogeographic Relationships  .........................................................................................................................................   77 Gordon McPherson

How Old are the Kauri (Agathis microphylla) Trees?  ...............................................................................................   83 Jonathan Palmer

The Flora of Santo ....................................................................................................................................................................   89 Some New, Characteristic or Remarkable Species ..........................................................................................    89 Gordon McPherson & Jérôme Munzinger

Focus on Araliaceae: Several Genera Exemplify Santo's Melanesian Biogeographic Relations .........................................    90 Porter P. Lowry II & Gregory M. Plunkett

Focus on Geissois (Cunoniaceae): Another Example of the Melanesian Connection ............................................................................................    93 Yohan Pillon

Focus on Pandans ................................................................................................................................................................    94 Thomas Haevermans

Focus on Orchids ..................................................................................................................................................................    97 Marc Pignal

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Contents

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Focus on Palms ......................................................................................................................................................................  102 Jean-Michel Dupuyoo

Focus on Ferns .......................................................................................................................................................................  105 Germinal Rouhan

Focus on Bryophytes ..........................................................................................................................................................  110 Elizabeth A. Brown

Fungi, the Forgotten Kingdom  ........................................................................................................................................  113 Bart Buyck

terrestrial fauna ...............................................................................................................................................................  117 coordinated by Bruno Corbara

IBISCA-Santo Biodiversity Along an Altitudinal Gradient  .......................................................................  119 Bruno Corbara on behalf of the IBISCA network

Insects on Santo ......................................................................................................................................................................... 123 Focus on Orthoptera ..........................................................................................................................................................  123 Laure Desutter-Grandcolas, Sylvain Hugel & Tony Robillard

Termites in Santo: Lessons from a Survey in the Penaoru Area .............................................................  128 Yves Roisin, Bruno Corbara, Thibaut Delsinne, Jérôme Orivel & Maurice Leponce

Focus on Bees and Wasps ..............................................................................................................................................  131 Claire Villemant

Myrmecophily in Santo: A Canopy Ant-Plant and its Expected and Less Expected Inhabitants 143 Bruno Corbara

Beetles in Saratsi Range, Santo ..................................................................................................................................  146 Alexey K. Tishechkin, Jürgen Schmidl

Lepidoptera in Vanuatu: Fauna, Geography and the IBISCA-Santo Project .....................................  155 Roger L. Kitching

Other Invertebrates .................................................................................................................................................................  161 Diversity of Spiders .............................................................................................................................................................  161 Christine Rollard

Some Arthropods as Expressed in the Words of Penaoru Villagers .....................................................  167 Bruno Corbara

Indigenous Land Snails .................................................................................................................................................... 169 Benoît Fontaine, Olivier Gargominy & Vincent Prié

The Vertebrates of Santo  .....................................................................................................................................................  179 Terrestrial Bird Communities  ........................................................................................................................................  179 Nicolas Barré, Thibaut Delsinne & Benoït Fontaine

Amphibians and Reptiles ................................................................................................................................................  187 Ivan Ineich

rivers and other freshwater habitats .....................................................................................................  237 coordinated by Philippe Keith

Freshwater Habitat Types  ..................................................................................................................................................  239 Philippe Keith & Clara Lord

Freshwater Biota .......................................................................................................................................................................  242 Focus on Fish, Shrimps and Crabs  ...........................................................................................................................  242 Philippe Keith, Clara Lord, Philippe Gerbeaux & Donna Kalfatak

Focus on Aquatic Insects  ...............................................................................................................................................  251 Arnold H. Staniczek

Focus on Freshwater Snails  ........................................................................................................................................... 257 Yasunori Kano, Elen E. Strong, Benoît Fontaine, Olivier Gargominy, Matthias Glaubrecht & Philippe Bouchet

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The Natural History of Santo

caves and soils ...................................................................................................................................................................... 265 coordinated by Louis Deharveng

The Karst Team ...........................................................................................................................................................................  267 Louis Deharveng & Anne-Marie Sémah

Karst and Caves .........................................................................................................................................................................  269 Bernard Lips, Franck Bréhier, Denis Wirrmann, Nadir Lasson, Stefan Eberhard, Josiane Lips & Louis Deharveng

Caves as Archives ...................................................................................................................................................................... 278 Denis Wirrmann, Jean-Christophe Galipaud, Anne-Marie Sémah & Tonyo Alcover,

Ni-Vanuatu Perception and Attitudes Vis-à-Vis the Karstic Environment  ................................  284 Florence Brunois

Karst Habitats of Santo .......................................................................................................................................................  288 Focus on Soils  .......................................................................................................................................................................  288 Anne Bedos, Vincent Prié & Louis Deharveng

Focus on Cave Terrestrial Habitats............................................................................................................................  296 Louis Deharveng, Anne Bedos, Vincent Prié & Éric Queinnec

Focus on Guano  ...................................................................................................................................................................  300 Louis Deharveng, Josiane Lips & Cahyo Rahmadi

Focus on Blue Holes  .........................................................................................................................................................  306 Stefan Eberhard, Nadir Lasson & Franck Bréhier

Focus on the Loren Cave  ................................................................................................................................................. 310 Franck Bréhier, Sephan Eberhard & Nadir Lasson

Focus on Anchialine Fauna  ........................................................................................................................................... 312 Geoff Boxshall & Damià Jaume

Karst Biota of Santo ................................................................................................................................................................ 316 Focus on Bats  ......................................................................................................................................................................... 316 Vincent Prié

Fish and Shrimps of Santo Karstic Systems  ......................................................................................................  323 Marc Pouilly & Philippe Keith

Focus on Springtails  ........................................................................................................................................................... 327 Louis Deharveng & Anne Bedos

Focus on Microcrustaceans  ..........................................................................................................................................  331 Damià Jaume, Geoff Boxshall & Eric Queinnec

Marine Ecosystems ............................................................................................................................................................ 335 coordinated by Philippe Bouchet

Benthic Algal and Seagrass Communities from Santo Island in Relation to Habitat Diversity ......................................................................................................................................... 337 Claude E. Payri

The Position of Santo in Relation to the Centre of Maximum Marine Biodiversity (the Coral Triangle) ..................................................................................................................................  369 Bert W. Hoeksema & Adriaan Gittenberger

Focus on Selected (Micro)Habitats ...........................................................................................................................  373 Sulfide Rich Environments .............................................................................................................................................  373 Yasunori Kano & Takuma Haga

Marine Interstitial ................................................................................................................................................................  375 Timea Neuser

Mangroves Environments of South East Santo .................................................................................................  377 Jean-Claude Plaziat & Pierre Lozouet

Focus on Selected Biota  .....................................................................................................................................................  383 Checklist of the Fishes ...................................................................................................................................................... 383 Ronald Fricke, John L. Earle, Richard L. Pyle & Bernard Séret

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Unusual and Spectacular Crustaceans ....................................................................................................................  410 Tim-Yam Chan, Masako Mitsuhashi, Charles H.J.M. Fransen, Régis Cleva, Swee Hee Tan, Jose Christopher Mendoza, Marivene Manuel-Santos & Peter K.L. Ng

The Marine Molluscs of Santo ..................................................................................................................................... 421 Philippe Bouchet, Virginie Héros, Pierre Lozouet, Philippe Maestrati & Rudo von Cosel

A Rapid Assessment of the Marine Molluscs of Southeastern Santo ................................................. 431 Fred E. Wells

Molluscs on Biogenic Substrates................................................................................................................................. 438 Anders Warén

Marine Partnerships in Santo's Reef Environments: Parasites, Commensals and Other Organisms that Live in Close Association ..............................  449 Stefano Schiaparelli, Charles Fransen & Marco Oliverio

Seaslugs: The Underwater Jewels of Santo ......................................................................................................... 458 Yolanda E. Camacho & Marta Pola

Man and nature ...................................................................................................................................................................  465 coordinated by Michel Pascal

Pre-European Times ...............................................................................................................................................................  467 Vertebrate Pre-Human Fauna of Santo: What Can we Expect to Find? .............................................. 467 Joseph Antoni Alcover

The Prehistory of Santo ...................................................................................................................................................  469 Jean-Christophe Galipaud

Introduced Biota ........................................................................................................................................................................ 476 Overview: Introduced Species, the "Good", the "Worrisome" and the "Bad" .................................  476 Michel Pascal, Olivier Lorvelec, Nicolas Barré, Michel de Garine-Wichatitsky & Marc Pignal

Focus on Synanthropic Mammals .............................................................................................................................  480 Olivier Lorvelec & Michel Pascal

Focus on Feral Mammals ...............................................................................................................................................  483 Michel de Garine-Wichatitsky & Anthony Harry

Focus on Alien Birds  .........................................................................................................................................................  488 Nicolas Barré

Focus on Introduced Amphibians and Reptiles  ..............................................................................................  490 Olivier Lorvelec & Michel Pascal

Focus on Introduced Fish  ..............................................................................................................................................  494 Philippe Keith, Clara Lord, Donna Kalfatak & Philippe Gerbeaux

Focus on Alien Land Snails  ..........................................................................................................................................  495 Olivier Gargominy, Benoît Fontaine & Vincent Prié

Endemic, Native, Alien or Cryptogenic? The Controversy of Santo Darkling Beetles (Insecta: Coleoptera: Tenebrionidae)  ...................  500 Laurent Soldati

The Case of Two Invasive Species: Mikania micrantha and Merremia peltata  .........................  503 Marc Pignal

Man Santo in his Environment .....................................................................................................................................  508 Food-Garden Biodiversity in Vanuatu  ....................................................................................................................  508 Sara Muller, Vincent Lebot & Annie Walter

At the Junction of Biological Cycles and Custom: the Night of the Palolo  .................................... 515 Laurent Palka

Ni-Vanuatu Perceptions and Attitudes Vis-à-Vis Biodiversité  ................................................................. 516 Florence Brunois & Marine Robillard

the santo 2006 expedition ......................................................................................................................................... 523 The Santo 2006 Expedition from an Ethnologist's Point of View ...................................................... 525 Elsa Faugère

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The "Making of" Santo 2006 ............................................................................................................................................. 529 Philippe Bouchet, Hervé Le Guyader, Olivier Pascal

Santo 2006 Expedition in the Classroom .............................................................................................................. 549 Sophie Pons & Alain Pothet

Santo 2006 Expedition Participants List ................................................................................................................ 550 Acknowledgements ................................................................................................................................................................. 553 Bibliography....................................................................................................................................................................................  557 Addresses of the Authors..................................................................................................................................................... 564

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in Bouchet P., Le guyader H. & pascal O. (Eds), The Natural History of Santo. MNHN, Paris; IRD, Marseille; PNI, Paris. 572 p. (Patrimoines naturels; 70).

Fungi

the Forgotten Kingdom Bart Buyck

Mushrooms are often looked upon as some kind of odd vegetable but thanks to technological progress it has now been clearly established that they belong to a very diverse group of organisms we call "fungi", which are neither plants nor animals. As a matter of fact, the fungi represent a separate kingdom of living organisms whose affinities are more with the animal world than with plants. Fungi are not studied by botanists, but by mycologists. Unfortunately, mycologists

are much harder to find than the fungi they study and we still know very little about the fungi on our planet. As past inventories focused principally on plant and animal biodiversity, we still know surprisingly little about fungal diversity. Scientists have so far named less than 5 % of the estimated minimum of 1.5 million species. It is, therefore, not surprising that we knew hardly anything about the fungi that are present on the island of Santo at the onset of this large inventory project. We have found no published records of past inventories of the fungi of Santo.

Fungi of Santo: a decomposer community

By just looking at what kind of animals and plants inhabit Santo, we may already have a hint of what groups of fungi to expect on the island. Indeed, one of the first things that may strike you when looking at the list of the forest trees that are present there, is the absence of those mushroom-associated tree families that dominate forest types elsewhere in the world: Santo lacks not only Fagaceae (oak, beech, chestnut, etc.), Betulaceae (birch, poplar, alder, etc.) and Pinaceae (pine, spruce or fir, etc.) that are so common in the northern hemisphere, but the island also has none of the locally dominant symbiotic trees from the southern hemisphere. Santo lacks for example eucalypts and some mushroomassociated Myrtaceae, common in Australia, nor is the island home to tree species of the predominantly Asian Dipterocarps or to some of Africa’s

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widely dispersed legumes (Isoberlinia, Brachystegia, Julbernardia, etc.) in the family Caesalpiniaceae. It is therefore not surprising that we did not encounter any of the above-mentioned, typical symbiotic forest mushrooms on Santo, although some fungal species may perhaps be associated on the island with the roots of the amphipacific legume tree Intsia bijuga, which was common on Santo but is under strong pressure from deforestation, or with the roots of coastal Pisonia (Nyctaginaceae). The absence of typical forest mushrooms does not by any means imply that you will encounter very few fungi when exploring Santo. Of course, the climatic conditions have to allow for their growth! Indeed, fungi need moisture for fruiting, and at least the soft fleshy mushrooms will only be present for a very limited

Photo B. Buyck

Fungi are generally depicted as organisms that thrive on the decomposition of dead organic material and wastes. Many fungal groups, however, are entirely composed of symbionts or parasites that are highly dependant for their development on specific groups of living plants or animals. The latter is also the case for many — probably even most — of the widely known, larger forest mushrooms that appeal so much to the general public. Both these typical forest mushrooms and their host trees can not survive without forming root symbiotic structures (called "ectomycorrhiza") in the soil. Examples of such symbiotic forest mushrooms include not only such reputed edibles as morels, truffles, chanterelles and boletes but also comprise several thousand species belonging to more common and very diverse mushroom genera such as Amanita, Russula, Lactarius, Cortinarius, Tricholoma, Hebeloma, Inocybe, etc.

Figure 134: Polypores are very common and diverse in Santo’s forests.

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Photo B. Buyck

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Photo B. Buyck

Vegetation and Flora

Figure 136: It is merely a matter of one or two hours for the delicate Dictyophora species too change from the egg-stage to their fully expanded mature form.

time of the year when rains are abundant. Mushroom hunting will therefore be more rewarding on the much wetter eastern part of Santo. The much tougher polypores are considerably less dependant on precipitation and may remain attached to logs and trunks of fallen or standing trees for many years and continue their growth with the arrival of every new rainy season. These large, tough polypores are the most prominent

representatives of the decomposer community that is so typical of Santo’s forests. One of the best places we visited was without any doubt the coastal dense forest reserve at Matantas. An overwhelming diversity of polypores in various colors and sizes was growing on almost every fallen log and branch on the ground (Fig. 134). Naturalists who have already looked for fungi in other tropical forests will easily recognize some common pantropical polypore genera such as Cymatoderma, Microporus, Pycnoporus and Polyporus. We particularly found the genus Polyporus itself to be strikingly diverse on Santo and we recorded close to fifteen different species during our short visit. One of these — an as yet undescribed species (Fig. 135) — is even locally consumed and bears a distinct resemblance to the European Polyporus squamosus.

Photo B. Buyck

Figure 137: Cookeina speciosa, by far the most common species of this genus, comes in various colours, from white over yellow to pinkish red.

Most of the decomposer fungi of Santo are smaller, ephemeral, fleshy species that remain present for only a few hours or perhaps a few days, the time needed to produce and disperse their spores. Examples among some of the more common, gilled mushroom genera include wood decomposers such as Pleurotus, Lentinus, Schizophyllum or the very beautiful yelloworange Cyptotrama asprata. Around the completely

Figure 138: Cookeina tricholoma is a close relative of C. speciosa and easily recognized by the long hairs that cover its outer surface.

Photo B. Buyck

Photo B. Buyck

Figure 135: This still undescribed Polyporus was found on dead tree trunks and was said to be edible and locally well-liked.

Figure 139: This strange, water-filled ascomycete is Galiella celebica, a pantropical species.

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....... decomposed and rotten remains of logs and truncs, species of Pluteus and Psathyrella form sometimes vast colonies of many dozens of fruit bodies. Such very nutrient-rich sites also favour the proliferation of spectacular Dictyophora species (Fig. 136). Also many ascomycetes decompose wood but most species are very small. Among some of the larger ascomycetes, we can cite the many bright yellow, pink, red or purple cups of the various species of the genus Cookeina (Figs 137 & 138) that sprout like colourful flowers from the smaller twigs and branches that have fallen on the forest floor, whereas another wood decomposing ascomycete, Galiella celebica (Fig. 139), keeps it


own water reserve safely stocked inside the cavity of its gelatinous fruitbody.

Among the leaf litter on the forest floor, many short-lived species of Coprinus, Lepiota, Marasmius and Mycena specialise in the decomposition of leaves and other parts of particular plant species. Outside the forest, in the pastures around the villages, slender species of Coprinus, Psilocybe, Panaeolus and other delicate mushrooms pop up everywhere after heavy rain, especially on or near cowdung or excrement left by horses, pigs and other animals.

Fungi that light up in the dark!

Photo B. Buyck

Bioluminescent fungi are not so rare as one might think. They occur all over the globe and have been known since ancient times. They produce light continuously but the human eye needs to wait for darkness before it can detect the light emission. In a very recent overview of bioluminescence in fungi, these organisms are placed in three different phylogenetic groups: the Omphalotus lineage, the Armillaria lineage and the Mycenoid lineage. All three comprise exclusively white spored basidiomycetes. The first two lineages are also common in France for example. The Jack-O-Lantern (Omphalotus olearius, "le pleurote de l’olivier" in French) is a good example of a medium to large-sized Mediterranean mushroom of the first lineage. The Armillaria lineage comprises some extremely common mushroom species that are

responsible for the "glowing wood" phenomenon — or "foxfire" as it has been known since ancient time — because it is not the mushroom itself, but the mycelium and mycelial cords inside the wood that emit light. On Santo, we observed only bioluminescent species of the Mycenoid lineage (Fig. 140). These are predominantly much smaller fungi, in this case species of Mycena and Filoboletus. Apart from the proven fact that the light attracts insects that may (perhaps) contribute to spore dispersal (in taxa whose fruit bodies are light emitting), there is no consensus about the "why" of this phenomenon. Whether it favours detoxification of some of the metabolites resulting from the wood degradation or is simply a by-product with no particular selective advantage, we do not know. The exploration of Santo opened a window on the incredibly diverse but still barely understood world of the fungi… and even though we will probably never be able to document the totality of the mycota of the quickly changing habitats of Santo, we can still profit from their ephemeral presence and enjoy their breath-taking beauty and ingenuity.

Photo B. Buyck

When night falls, the dancing lights of fire flies are a familiar sight for many travellers, but mushrooms that light up in the dark remain a surprising experience. Going out for a night walk in the forest can definitely be recommended: bioluminescent fungi are common on Santo and well known by the local population, who use them as a light source.

Figure 140: Bioluminiscent species of Mycena and Filoboletus are quite common in the wetter parts of Santo’s forests. Here the same specimens of a Mycena photographed during daylight and at night.

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Santo The islands of the Pacific are renowned for the high levels of endemism of, and threats to, their unique faunas and floras. Espiritu Santo, affectionately known simply as Santo, is an island of superlatives: the largest and highest in Vanuatu, Santo is an extraordinary geographical and cultural microcosm, combining reefs, caves, mountains, satellite islands, and a history of human habitation going back 3 000 years. In the spirit of famous voyages of discovery of the past, the Santo 2006 expedition brought together over 150 scientists, volunteers and students originating from 25 countries. With contributions by more than 100 authors, The Natural History of Santo is a lavishly illustrated homage to the biodiversity of this "planet-island". Bridging the gap between scientific knowledge and conservation and education, The Natural History of Santo was written with local stakeholders as well as armchair naturalists from all over the world in mind.

Les îles du Pacifique sont célèbres pour le très haut niveau d’endémisme et la grande vulnérabilité de leurs faunes et de leurs flores. L'île d'Espiritu Santo, ou Santo, cumule les superlatifs : la plus grande et la plus haute du Vanuatu, Santo est un extraordinaire microcosme géographique et culturel, avec récifs, grottes, montagnes, îles et îlots satellites, et une occupation humaine qui remonte à 3 000 ans. Renouant avec l'esprit des "Grandes Expéditions Naturalistes", l’expédition Santo 2006 avait mobilisé sur le terrain plus de 150 scientifiques, bénévoles et étudiants de 25 pays. Petit tour de force éditorial avec plus de 100 auteurs, ce Natural History of Santo est un éloge de la biodiversité de cette "île-planète". À la fois beau livre richement illustré et bilan des connaissances scientifiques, The Natural History of Santo se veut un outil de connaissance pour sa conservation durable. Il s'adresse autant aux acteurs locaux du développement et de l'éducation qu'aux naturalistes du monde entier.

ISSN 1281-6213 ISBN MNHN : 978-2-85653-627-8 ISBN IRD : 978-2-7099-1708-7

9 782856 536278

Prix : 59  TTC